A new species of Dendrophidion (Serpentes: Colubridae) from the Atlantic Rain Forest of Northeastern Brazil

Abstract

A new species of Dendrophidion belonging to the D. dendrophis species group is described from Mata do Engenho Coimbra (08°59’S, 35°53’W; 526 m above sea level), Municipality of Ibateguara, in the Atlantic Rainforest remnants of the State of Alagoas, northeastern Brazil. Dendrophidion atlantica sp. nov. is characterized by having 154–163 ventral scales, 140–160 subcaudal scales, tail length 62.2–74.8% of snout–vent length, collar absent, head uniformly brown and dorsal ground color brown, paler on anterior third, with cream transversal lines (one half a scale long), bordered anteriorly and posteriorly by dark brown lines (one half a scale long), distributed from the neck to the tail; hemipenis single, subcylindrical, unicapitate, and unicalyculate; calyces large, well defined, papillate; a series of 12 large spines just
below the capitulum, on the asulcate and lateral sides; a series of four spines, two large laterals and two small between them, in the basal region of the asulcate side of the organ body; two large spines on the lateral distal areas of the sulcate side of the hemipenial body; sulcus spermaticus centrolineal, bifurcating at the tip of the capitulum.

Full text

62 Accepted by D. Gower: 24 Nov. 2010; published: 10 Dec. 2010
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2010 · Magnolia Press
Zootaxa 2719: 62–68 (2010)
www.mapress.com/zootaxa/Article
A new species of Dendrophidion (Serpentes: Colubridae) from the Atlantic Rain
Forest of Northeastern Brazil
ELIZA MARIA XAVIER FREIRE1,4, ULISSES CARAMASCHI2 & UBIRATAN GONÇALVES3
1Departamento de Botânica, Ecologia e Zoologia, Centro de Biociências, Universidade Federal do Rio Grande do Norte, Campus
Universitário, 59072-970 Natal, Rio Grande do Norte, Brasil. E-mail: elizajuju@ufrnet.br
2Universidade Federal do Rio de Janeiro, Museu Nacional, Departamento de Vertebrados, Quinta da Boa Vista, 20940-040 Rio de
Janeiro, Rio de Janeiro, Brasil. E-mail: ulisses@acd.ufrj.br
3Museu de História Natural, Universidade Federal de Alagoas, 57000-000 Maceió, Alagoas, Brasil. E-mail: ugsbogertia@gmail.com
4Corresponding author
Abstract
A new species of Dendrophidion belonging to the D. dendrophis species group is described from Mata do Engenho
Coimbra (08°59’S, 35°53’W; 526 m above sea level), Municipality of Ibateguara, in the Atlantic Rainforest remnants of
the State of Alagoas, northeastern Brazil. Dendrophidion atlantica sp. nov. is characterized by having 154–163 ventral
scales, 140–160 subcaudal scales, tail length 62.2–74.8% of snout–vent length, collar absent, head uniformly brown and
dorsal ground color brown, paler on anterior third, with cream transversal lines (one half a scale long), bordered
anteriorly and posteriorly by dark brown lines (one half a scale long), distributed from the neck to the tail; hemipenis
single, subcylindrical, unicapitate, and unicalyculate; calyces large, well defined, papillate; a series of 12 large spines just
below the capitulum, on the asulcate and lateral sides; a series of four spines, two large laterals and two small between
them, in the basal region of the asulcate side of the organ body; two large spines on the lateral distal areas of the sulcate
side of the hemipenial body; sulcus spermaticus centrolineal, bifurcating at the tip of the capitulum.
Key words: Serpentes, Colubridae, Dendrophidion atlantica sp. nov., Atlantic Rain Forest, taxonomy
Introduction
The colubrid genus Dendrophidion Fitzinger, 1843 consists of a group of eight described species of racer–like
snakes distributed from southern Mexico to northern Brazil and Bolivia (Lieb 1988). These eight species were
placed into three species groups, including the dendrophis and percarinatum groups, as well as a single-species
incertae sedis group (Lieb 1988). Lieb (1988) placed three species [D. dendrophis (Schlegel, 1837), D. nuchale
(Peters, 1863), and D. vinitor Smith, 1941] in the dendrophis group. This group occurs throughout the region
occupied by the entire genus, with D. vinitor, the northernmost species occurring from southern Mexico to western
Colombia, D. nuchale ranging from Belize and central Guatemala to northwestern Ecuador, with disjunct
populations in Venezuela, and the southernmost species D. dendrophis inhabiting the northern Amazonian region
from eastern Ecuador and Peru through northern Brazil to the Guianas (Lieb 1988; Köhler 2008). Lieb (1988)
allocated four species [D. bivittatum (Duméril, Bibron and Duméril, 1854), D. brunneus (Günther, 1858), D.
paucicarinatum (Cope, 1894), and D. percarinatum (Cope, 1893)] to the percarinatum group. Collectively, these
species range from northern Honduras to southwestern Ecuador (McCranie et al. 2006). Lieb (1988) placed a
single species (D. boshelli Dunn, 1944) in his incertae sedis group, which is distributed in central Colombia.
Lieb (1988) also recognized an “undescribed complex of populations in the southern Amazon Basin and
adjacent areas” as related to D. dendrophis and D. nuchale, indicating that the status of these populations was “the
subject of work in progress.” Inasmuch as this work appears to have never been completed, we take this
opportunity to describe as a new species one of the members of this complex from the Atlantic Rainforest remnants
of northeastern Brazil.

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A NEW DENDROPHIDION FROM NORTHEASTERN BRAZIL
Material and methods
Specimens examined are deposited in the Museu Nacional, Rio de Janeiro, Rio de Janeiro, Brazil (MNRJ), Museu
de História Natural, Universidade Federal de Alagoas, Maceió, Alagoas, Brazil (MUFAL), Museu de História
Natural do Capão da Imbuia, Curitiba, Paraná, Brazil (MHNCI), and Coleção Herpetológica of the Departamento
de Botânica, Ecologia e Zoologia, Universidade Federal do Rio Grande do Norte, Natal, Rio Grande do Norte,
Brazil (CHBEZ); specimens examined are indicated in the Appendix 1.
Terminology for cephalic shields follows Lieb (1988) and ventrals were counted following Dowling (1951).
Terminology for the hemipenis follows Dowling and Savage (1960), as augmented by Myers and Campbell (1981)
and Zaher (1999). Hemipenes were examined in the everted condition; when necessary they were everted using the
techniques described by Pesantes (1994). Sex was determined by the presence or absence of everted hemipenes or
through a ventral incision at the base of the tail. Measurements were taken with calipers to the nearest 0.1 mm
under a binocular stereomicroscope, except for snout–vent length (SVL) and tail length (TL), which were taken
with a flexible ruler to the nearest 1 mm.
Dendrophidion atlantica sp. nov.
“Atlantic Forest Racer”
(Figures 1–3)
Dendrophidion dendrophis—Freire 1999.
Holotype: MNRJ 17018, adult male (Fig. 1), from Mata do Engenho Coimbra (08°59’S, 35°53’W; 526 m above
sea level), Municipality of Ibateguara, State of Alagoas, Northeastern Brazil, collected by Ubiratan Gonçalves on
05 December 2006.
FIGURE 1. General view in life of the holotype of Dendrophidion atlantica sp. nov. (MNRJ 17018; SVL 595 mm).
Paratypes: BRAZIL: ALAGOAS: Mata do Catolé (09°40’S, 35°43’W; 122 m altitude), Municipality of
Maceió: CHBEZ 853, female, collected by E. M. X. Freire on 29 September 2002; CHBEZ 697, female, collected

FREIRE ET AL.64 · Zootaxa 2719 © 2010 Magnolia Press
by E. M. X. Freire on 24 May 2003; CHBEZ 2201, female, MUFAL 6064, male, collected by E. M. X. Freire on 05
May 1995; MUFAL 6065, male, collected by E. M. X. Freire on 24 de October 1996; MNRJ 17020, male, collected
by E. M. X. Freire on 02 November 2001. Mata da Salva (09°32’S, 35°49’W; 85 m altitude), Municipality of Rio
Largo, MUFAL 327, female, collected by E. M. X. Freire on 13 September 1992; CHBEZ 2202, male, collected by
E. M. X. Freire on 01 November 1993. Mata do Rio Messias, Municipality of Messias (09°26’S, 35°47’W; 81 m
altitude), MNRJ 17021, male, collected by J. Luiz on 01 May 1993. Mata da Bananeira (09o14’S, 35o48’W; 500 m
altitude), Municipality of Murici, MNRJ 17019, female, collected by E. M. X. Freire on 04 November 1994;
MUFAL 2245, male, collected by E. M. X. Freire on 22 November 1995.
FIGURE 2. Lateral (A), dorsal (B), and ventral (C) views of the head of Dendrophidion atlantica sp. nov. (holotype,
MNRJ 17018). Bar equals 10 mm.

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A NEW DENDROPHIDION FROM NORTHEASTERN BRAZIL
FIGURE 3. Sulcate (A) and asulcate (B) views of the hemipenis of Dendrophidion atlantica sp. nov. (holotype, MNRJ
17018). Bar equals 5 mm.
Diagnosis: A species belonging to the Dendrophidion dendrophis species group, based on the presence of: (1)
17 anterior dorsal scale rows; (2) all dorsal scale rows strongly keeled; (3) dorsocaudal scale row reduction from
eight to six occurring posterior to the 25th subcaudal; and (4) hemipenes with enlarged basal spines. This species is
distinguished from the other members of this species group (D. dendrophis, D. nuchale, and D. vinitor) by having
the following features: (1) 154–163 ventral scales; (2) 140–160 subcaudal scales; (3) TL 62.2–74.8% SVL; (4)
collar absent; (5) head uniformly brown, dorsum brown, paler on anterior third, with transverse middorsal and
lateral cream markings (one half a scale long), located within a dark brown transverse band about one and one-half
a scale long at midbody; (6) hemipenis simple, sybcylindrical, unicapitate, and unicalyculate, calyces large, well
defined, papillate, with a set of 12 large spines just proximate to the capitulum on the asulcate side, and a
centrolineal sulcus spermaticus bifurcating at the distal tip of the capitulum; (7) distribution in remnants of the
Atlantic Rainforest in northeastern Brazil.
Description of the holotype: Adult male, SVL 595 mm, TL 445 mm (74.8% of SVL); head length 22.2 mm
(3.7% of SVL); body diameter 12.5 mm (2.1% of SVL); head width 11.7 mm (52.7% of head length); interorbital
distance 9.8 mm; rostro-orbital distance 6.5 mm (66.3% interorbital distance); naso-orbital distance 5.1 mm; head
distinct from body, slightly arched in lateral view, elliptical in dorsal view; snout truncate in lateral and dorsal
views (Fig. 2 A–B); rostral semilunar, barely visible in dorsal view; internasal 3.1 mm long, 3.5 mm wide;
prefrontal 3.2 mm long, 3.5 mm wide; supraocular elongate in dorsal view, 6.4 mm long, 3.1 mm wide; frontal

FREIRE ET AL.66 · Zootaxa 2719 © 2010 Magnolia Press
subtriangular in dorsal view, 6.8 mm long, 2.4 mm wide; parietal 7.7 mm long, 5.2 mm wide; nasal divided; nostril
located between prenasal and postnasal; prenasal twice higher than long; postnasal similar in height to prenasal,
slightly longer than high; loreal 2.5 mm long, 1.6 mm high; eyes large, eye diameter 4.2 mm, pupil round; two
preoculars, the upper 3.0 mm high, 2.1 mm wide, distinctly larger than lower; two postoculars; upper postocular
taller (2.6 mm) and longer (1.1 mm) than lower postocular; temporals 2+2; anterior upper temporal 2.4 mm long,
1.3 mm high; first inferior temporal 5.2 mm long, 1.8 mm high; upper posterior temporal elongate, 3.7 mm long,
1.2 mm wide; inferior posterior temporal 2.6 mm long, 1.3 mm wide; 9–9 supralabials, fourth to sixth contacting
the eye; mental triangular, 3.0 mm width, broader than long (Fig. 2 C); 9–9 infralabials, first to fifth contacting the
anterior chinshield, posterior part of fifth and sixth contacting the posterior chinshield; first pair of infralabials in
contact behind mental, preventing contact between mental and chinshields; anterior chinshields 4.6 mm long, 1.8
mm wide; posterior chinshields 7.6 mm long, 2.1 mm wide; one pair of gular scales; two preventrals. Dorsal scales
carinate, with paired apical pits on some of them; dorsals in 17–17–15 rows; 154 ventrals; cloacal scute single; 152
divided subcaudals; dorsocaudal scale reduction from eight to six posterior to subcaudal 32.
In life (Fig. 1), head uniformly brown; collar absent; dorsum brown, paler on anterior third, with transverse
cream markings one half a scale long, bordered anteriorly and posteriorly by dark brown lines one half scale long,
distributed from the neck to the tail; in some parts of dorsum, these lines are interrupted in the middle and the
patterned scales of each lateral line are disposed alternately; infralabials cream; ventral scutes cream with lateral
margins dark brown due to the extension of the dorsal color. In preservative, the dorsal color is brownish gray with
transverse markings white bordered by black and the venter is cream.
Retracted hemipenis bifurcates and extends to the level of the eighth subcaudal scale. Everted organ (Fig. 3 A–
B) single, subcylindrical, unicapitate, and unicalyculate; capitular groove distinct on both sides of the organ;
capitulum about one third of the length of the entire organ; calyces large, well defined, papillate; a group of 12
large spines just below the capitulum, on the asulcate and on the lateral side; a group of small spines on the distal
portion and a series of four spines, two large laterals and two small between them, in the basal region of the
asulcate side of the organ body; on the sulcate side, the organ body presents medium sized spines just below the
capitulum and two large spines on the lateral distal areas; sulcus spermaticus deep, with margins thick, smooth;
sulcus spermaticus centrolineal, bifurcating at the tip of the capitulum and each small branch ending in small re-
entrances.
Variation among paratypes: There is little variation among the paratypes, primarily in some scales counts.
There are 9–9 or 9–10 supralabials, 9–9 or 9–10 infralabials, 154–157 ventrals in males and 157–163 in females,
and 140–160 subcaudals in males and 145–153 in females. Maximum total length (SVL plus TL) is 1162 mm in
males and 1024 mm in females.
Comparison with other members of the Dendrophidion dendrophis group: Dendrophidion atlantica can be
distinguished from D. dendrophis (characteristics of D. atlantica followed by those of D. dendrophis, in
parentheses) by having a lower number of subcaudal scales (140–160 vs. 171–195; Lieb 1988), a shorter tail (mean
TL 40.4% of total length in males and 41.2% in females vs. 44.9% in males and 45.1% in females; Prudente et al.
2007), narrower transverse white markings and dark brown crossbands (one half and one and one half a scale long,
respectively, vs. about one and two scales long, respectively; Lieb 1988), hemipenis unicalyculate vs. flounced,
with a group of 12 large spines just proximate to the capitulum vs. 10 spines, and a group of four spines (two large
lateral spines with two smaller ones between them) on the basal portion of the sulcate side of the hemipenial body
vs. a group of three spines (two very large lateral spines with one slightly smaller one between them).
Dendrophidion atlantica is distinguished from D. nuchale by having white and dark brown markings the length of
the body vs. having the anterior third of the body paler and patternless and the posterior third darker and patterned
usually with rows of pale yellow ocelli (Lieb 1988) and a hemipenis with small basal spines vs. two to four large
basal spines (Lieb 1988). Dendrophidion atlantica is differentiated from D. vinitor by having a higher number of
subcaudal scales (140–160 vs. 111–128; Lieb 1988), narrow pale markings on the body vs. broad pale crossbands
on the anterior portion of the body, and a hemipenis with basal spines vs. elongate, hooked villi on the proximal
portion; Lieb 1988).
Habitat and habits: We observed and/or collected 41 specimens of D. atlantica sp. nov., 35 of them inside
forest and six on its border. All specimens were found on the litter, except one observed descending from a small
tree to the ground. Inside the forest or on the border, the specimens always were found in the shade during the day
between 07h30min to 17h50min, with a peak of activity between 10h45min and 14h09min.

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A NEW DENDROPHIDION FROM NORTHEASTERN BRAZIL
All specimens, when located, remained motionless. When approaching to collect them, they made rapid
movements, giving the impression that they were moving far away, but actually the snakes had moved only one or
two meters ahead, becoming motionless again. When captured, the snake contorted the body and discharged
cloacal material. Seven specimens autotomised their tails during capture, and five succeeded in escaping using this
defense mechanism.
Geographical distribution: Known from five localities in the State of Alagoas, Brazil, all associated with the
Atlantic Rain Forest biome (Fig. 4).
Etymology: The specific name, used as a noun in apposition, is an allusion to the disjunct occurrence of this
new species in the Atlantic Rain Forest.
FIGURE 4. Distribution of Dendrophidion atlantica sp. nov. in the State of Alagoas, Brazil. 1: Mata do Catolé, Maceió;
2: Mata da Salva, Rio Largo; 3: Mata do Rio Messias, Messias; 4: Mata da Bananeira, Murici; 5: Mata do Engenho
Coimbra, Ibateguara (type locality).
Discussion
The nine species now comprising the genus Dendrophidion are distributed throughout the countries of
Mesoamerica (except for El Salvador) and the South American countries of Colombia, Ecuador, Peru, Bolivia,
Venezuela, all three of the Guianas, and Brazil (Lieb 1988; Uetz et al. 2009). Prior to the present work, the only
species known to occur in Brazil was D. dendrophis, distributed in the Amazonian region in the states of
Amazonas, Pará, Amapá, Maranhão, Acre, and Rondnia (Prudente et al. 2007). Dendrophidion dendrophis is
terrestrial to semi–arboreal and inhabits primary and secondary growth forests (Cunha and Nascimento 1993;
Martins and Oliveira 1998). The presence of a population in northeastern Brazil was reported by Lieb (1988), who
thought it possibly referable to a new species, but he did not document any specimens he examined. The habitat
and habits of D. atlantica are similar to those of D. dendrophis and we think it likely that the new species will
prove to feed on small, terrestrial anurans.
In the key to the species of Dendrophidion published by Lieb (1988), D. atlantica keys out to D. dendrophis,
but they can readily be distinguished on the basis of the numbers of subcaudal scales.

FREIRE ET AL.68 · Zootaxa 2719 © 2010 Magnolia Press
Acknowledgments
We deeply acknowledge Larry D. Wilson for his critical constructive review which substantially improved the
manuscript. We also thank the owners of the Usina Serra Grande, especially Mr. Clodoaldo Bakker, for their
hospitality and for allowing collecting in the Mata do Engenho, Ibateguara (AL); Selma Torquato for collecting
help in the field; Júlio César de Moura-Leite, Gilberto Alves de Souza Filho (MHNCI), and Gabriel Skuk
(MUFAL) for lending specimens under their care for study; Paulo Passos for hemipenial preparations and critically
reading an earlier draft; Paulo Roberto Nascimento for the hemipenial drawings; IBAMA/AL for logistical
support; and the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) for financial support.
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APPENDIX 1. Additional specimens examined.
Dendrophidion dendrophis—BRAZIL: AMAZONAS: Benjamin Constant, Rio Javari (MNRJ 498); Rio Itacoa (MNRJ
3025). PAR: Oriximin, Porto Trombetas, Sarac-Taquera (MNRJ 14923, 16806); Parauapebas, Serra dos Carajás
(MNRJ 19507, 19528). RONDÔNIA: Espigão d’Oeste, Fazenda Jaburi (MHNCI 9608). MATO GROSSO: (MNRJ
363); Rio Jauru, acima da Pedra Branca (MNRJ 362).